1. Department of Biological and Environmental Science, University of Jyväskylä, P.O. Box 35, 40014 University of Jyväskylä, Finland.
2. Department of Biology, University of Oulu, P.O. Box 3000, Finnish Environment Institute, 90014 Oulu, Finland.
3. Oulanka Research Station, Thule Institute, 90014 University of Oulu, Finland.